Instructors:                            Dr. Matthew J. Gray (mgray11@utk.edu)

Dr. Jason Hoverman (jhoverma@utk.edu)


Graduate Assistant:              Nathan Haislip (nhaislip@utk.edu)


Phone:                                    974-2740 (MG), 974-0831 (JT), 974-3897 (NH)

Office:                                    247 Ellington PSB (MG), 230 Ellington PSB (JT), 201 Ellington PSB (NH)

Meeting Time and Place     5:05–7:05 pm  T,R      113 PBB (2 field trips required: 12 March and 18 April)


Course Goal:              To expose students to the life history, diversity, ecology, conservation, and management of amphibians through a combination of lectures, readings, class discussions, labs, and field experiences.


Expected Outcomes:             Students that successfully complete WFS 433/533 will have a basic understanding of amphibian identification (larvae and adults), anatomy, life history, and ecology.  They will be aware of potential mechanisms of amphibian declines, and understand how to identify, collect and ship diseased amphibians to a diagnostic lab.  Students will understand how to sample amphibian populations and be aware of conservation strategies.  


Required Text:          The Ecology and Behavior of Amphibians, 2007 (ISBN 9780226893341)

Author:                      Kentwood D. Wells

Journal Papers:         Occasionally journal papers will be assigned instead of or to supplement the required text.  Papers will be provided in class or on the course website.



Academic Assessment:


Weights of Academic Assessments:

WFS 433


WFS 533

● Test #1        



● Test #1        


● Test #2



● Test #2


● Amphibian ID Exam



● Amphibian ID Exam







● Participation3,4



● Participation3,4


1Mini-presentations will be 12-17 minute persuasive presentations attempting to convince the audience of a specific cause for amphibian declines. 

2Graduate student lectures will be 45-55 minute presentations on an amphibian topic approved by Drs. Gray and Hoverman. 

3Participation includes attendance on the 2 field trips.  You will earn 5% for attending each field trip.

4If you miss a field trip, you can either: (1) attend the SEPARC meeting, (2) write a 10-page scientific paper on an amphibian topic of your choice, or (3) accept a 5% deduction in your final grade.


Your course grade will be determined using the following scale:



Final Weighted Percent



Final Weighted Percent






















Extra Credit:            


You can positively influence your final grade as much as 4.5% by volunteering for extra credit.  Volunteer work must be related to herpetofauna, and can include work on university projects or with government agencies or NGOs.  For every 8 hours of volunteer work, your final grade will be increased by 1.5% up to 4.5% (24 hours total).  All volunteering must be completed by 28 April 2009.  A volunteer form (see website) must be filled out by the supervising individual.  Scott Dykes and Pete Wyatt (TWRA non-game biologists) often are looking for volunteer assistance (Region 4 Office: 1-800-332-0900; scott.dykes@state.tn.us, pete.wyatt@state.tn.us).  Also, Wayne Schacher (consulting biologist for Seven Islands Wildlife Refuge) frequently needs help with herptofaunal surveys (457-4355 home, whschacher@natreserv.com).  Nathan Haislip (TA for WFS 433/533) will need assistance collecting tadpoles from farm ponds in January and April. 


Extra Credit Form


Extra credit also can be earned by attending the Annual Meeting of the Southeast Partners in Amphibian and Reptile Conservation: http://www.separc.org/.  The meeting is 19 – 22 February in Montreat, North Carolina.  Your final grade will be increased by 3.5% for attending the entire meeting.  Transportation will be provided.  If interested, you need to sign up by 22 January and register by 3 February if you are planning to travel with Drs. Gray and Hoverman.


Full Syllabus


Teaching Resources:


Required Tennessee Anurans

Required Tennessee Salamanders

Anuran Sonograms

Random Generation of Mini-Presentations

Tadpole Morphology (Haislip Lecture)

Practice Exam – TN Amphibian Identification


Required Readings:


TEST #1 Material


1)      Amphibian Diversity and Evolution

Required Readings:

Wells: pp. 1-15, 41-58, 65-74, 77-80

Supplemental Readings

Wells: pp. 16-41, 59-65, 75-77


2)      Amphibian Declines

Required Readings:

a.       What makes amphibians especially vulnerable to declines?

Wells: pp. 787-792

b.      What are some species in North America with relic populations?

Wells: pp: 793-794

c.       What is the region of the United States with the greatest number of species declining?  Also, be able to list a few species with distributions east of the Mississippi that are declining.

Wells: pp: 800-803

d.      Know which island in the South Pacific likely has the highest diversity of amphibian species per unit area in the world.

Wells: p. 795

e.       Be able to provide an argument for why we should care that amphibians are declining.

Wells: pp. 850-853

Supplemental Readings

Wells: pp. 816-850 (hypotheses for declines)


3)      Reproductive Strategies

Required Readings:

a.       Be able to describe different strategies for anuran oviposition in standing water.

Wells: pp. 465-468

b.      Be able to describe the differences between bubble and foam nests used in anuran oviposition and their adaptive significance.

Wells: pp: 472-478

c.       Be able to describe the differences between two modes of salamander reproduction and their adaptive significance.

Wells: pp. 488-493

d.      Be able to provide an explanation why some salamanders that deposit eggs in still water lay their eggs in clumps while others scatter single eggs.

Wells: pp. 788-489

e.       Know which family of salamanders only exhibits direct development.

Wells: p. 491

f.       Know the two salamander genera that exhibit ovoviviparity and viviparity.

Wells: p. 493

Supplemental Readings

Wells: pp. 465-493 (modes of anuran reproduction)

Zug Handout (pp. 169-189, Zug 1993, Herpetology, Academic Press).


4)      Amphibian Movements, Courtship, and Mating

Required Readings:

a.       Understand the difference between home range, migration and dispersal.

Wells: pp. 230-231

b.      Know the median home range for anurans and salamanders, and how they compare with birds and mammals.

Wells: pp: 230-231

c.       Know which age class disperses most frequently in an amphibian population.

Wells: pp. 243-244

d.      Be able to provide some hypotheses for the adaptive significance of homing (i.e., site fidelity). 

Wells: p. 253

e.       Know the principal site of extraocular photoreception and how amphibians use polarized light to orienteer.

Wells: pp. 261-264

f.       Be able to describe the 2 auditory channels in amphibians, and know which is sensitive to low frequencies (<300 Hz). 

Wells: pp. 322-324

g.      Be able to provide a couple examples of anurans that do not have vocal sacs, and an explanation of why this may have occurred.

Wells: p. 277

h.       Be able to describe unison bout calling and be able to provide some explanations why it may occur.

Wells: p. 297

i.        Schwartz (1991) proposed 3 hypotheses for the duration of unison bouts.  Be able to describe those and know which is most plausible.

Wells: p. 297

j.        Understand the difference between explosive vs. prolonged breeders, and be able to provide some reasons why cold-weather breeders and species that inhabit xeric environments may breed explosively.

Wells: pp. 339-341

k.      Understand factors that influence sexual selection in prolonged vs. explosive breeding systems.  Also, understand how energy allocation differs between prolonged and explosive breeders, and a few strategies that prolonged breeders may use to reduce energy spent on calling.

Wells: pp. 342-343

l.         Know the 4 ways that salamanders communicate, and be able to describe their primary function(s). 

Wells: pp. 404-411

m.    Know the 3 locations of chemical receptors in salamanders, and the 2 chemosensory organs in the nasal cavity.  Also, know which sex the chemosensory organs are usually larger and why this may occur.

Wells: pp. 417-418

n.      In the work performed by Robert Jaeger and Alicia Mathis on red-backed salamander territoriality, know the most important determinant of territory quality.  

Wells: pp. 424-425

o.      Be able to provide at least 2 explanations why inguinal amplexus is considered more primitive than axillary amplexus.

Wells: pp. 454-456

p.      Be able to describe the adaptive significance of internal fertilization via a spermatophore, and the difference between the duration that sperm survive in the spermatheca in ambystomatids vs plethodontids. 

Wells: pp. 459-461

q.      Understand the relationships between female body size and clutch size, female body size and egg size, egg size and clutch size, egg size and hatchling size, egg size and development rate, and developmental rate and temperature.

Wells: pp. 497-500

r.        Be able to describe selective advantages of species that produce small vs. large eggs, and the environmental constraints that may drive these relationships. 

Wells: pp: 502-504

s.       Know which mode of development has species that produce the largest eggs relative to body size.

Wells: p: 504

Supplemental Readings

Wells: pp. 254-266 (orienteering)

Wells: pp. 269-304, 314-316 (anuran calls)

                                    Wells: pp. 452-461 (external vs. internal fertilization)

Types of Anuran Calls: Types of Anuran Calls: MP3 file from The Calls of Frogs and Toads by Lang Elliot (Stackpole Books)


5)      Tadpole Development and Metamorphosis

Required Readings:

            What do tadpoles eat?  Altig et al. (2007)


a.       Understand how limb development is different between larval salamanders and tadpoles.

Wells: p. 559

b.      Be able to describe a few oral adaptations of larval salamanders that assist in suction feeding.

Wells: pp: 561-562

c.       Know whether salamander larvae are herbivorous, omnivorous or carnivorous.

Wells: p. 562

d.      Be able to explain some of the costs and benefits of cannibalism for larval salamanders.  What are some explanations for cannibalizing different species?

Wells: pp. 563-564

e.       Be able to provide some examples of tadpole species that delay lung development and why this may occur.

Wells: pp. 565

f.       Be able to provide some explanations why the limbs of tadpoles emerge late in development compared to salamanders.

Wells: pp. 565-566

g.      Understand the mechanism that induces carnivory in spadefoots found in the southwestern United States.  Also, understand how frequency of cannibalistic and omnivorous phenotypes differs between spadefoot species (S. bombifrons and S. multiplicata) and why this may occur. 

Wells: pp. 575-576

h.      Although most amphibian larvae are solitary, be able to provide some benefits of aggregation behavior and what may be the adaptive significance of each benefit. 

Wells: pp. 588-590

i.        Be able to describe the “selfish herd” effect and provide an example.

Wells: p. 589

j.        Be able to describe kin recognition and offer some explanations why this may be advantageous.

Wells: pp. 591, 593-596

k.      Be able to explain the factors that drive a larval amphibian to decide when to initiate metamorphosis according to Werner (1986, 1988) and Rowe and Ludwig (1986, 1988).  NOTE:  Your explanation should discuss differential mortality and growth rates in the aquatic and terrestrial environments and the amount of time between metamorphosis and the age of first reproduction.

Wells: pp. 601-602

l.        Know what hormone likely is responsible for initiating metamorphosis in response to an environmental stressor.

Wells: p. 608

Supplemental Readings

            Tadpoles: Macrophagous Predators Petranka and Kennedy 1999: Tadpoles: Macrophagous Predators

                                    Wells: pp. 564-566, 604-608


TEST #2 Material


1)      Phenotypic Plasticity

Required readings:   Study questions were sent by Dr. Hoverman in an email.

·         Wells pp. 601-603, 609-610, 618-628, 632-642

·         Gotthard and Nylin (1995): Oikos 74:3-17

·         Relyea (2007): Oikos 152:389-400

Supplemental readings:

·         Wells pp. 563-564, 573, 575, 596-597, 693-728


2)      Community Ecology



Required readings:   Study questions were sent by Dr. Hoverman in an email.

·         Wells pp. 694-696, 754-758, 768-778

·         Wellborn et al. (1996): Annual Review of Ecology and Systematics 27:337-363

·         Werner et al. (2007): Oikos 116:1697-1712

Supplemental readings:

·         Wells pp. 729-740, 779-783



Required readings:   Study questions were sent by Nathan in an email.

·         Impact of predation on amphibian populations (Wells pp. 681-693)

·         Defensive adaptations of adults (Wells pp. 709-716)

·         Aposematic coloration and mimicry (Wells pp. 721-727)

·         Competition and predation in larval salamander communities (Wells pp. 758-762)

·         Southerland 1986

Supplemental readings:

·         Amphibian predators (Wells pp. 645-681)

·         Predatory defenses (Wells pp. 715-727)

·         Chapter 15 introduction (Wells pp. 729-734)

·         Terrestrial competition (Wells pp. 745-753)


3)      Amphibian Sampling

Required readings:  

·         TAMP Protocol

·         Burton et al. (2007): Duration of Call Surveys


4)      Disease Sampling

Required readings:   Study questions were sent by Nathan in an email.

·         Shipping Protocol SOP (Gray Lab)

·         SE PARC Collection and Shipping Information Sheet (Miller and Gray)

·         Chapter 26 (sections 26.3 & 26.4): Disease Monitoring and Biosecurity (Green et al. 2009: in press): chapter emailed

Supplemental readings:

·         Chapter 26 (remaining sections): Disease Monitoring and Biosecurity (Green et al. 2009: in press): chapter emailed


5)      Estivation & Hibernation

Required readings:   Study questions were sent by email.

·         Wells pp. 94-98, 101, 120, 149-155, 188

Supplemental readings:

1.      Churchill and Storey (2005):  Dehydration tolerance in wood frogs


6)      Landscape Ecology

Supplemental readings:

1.      Joly et al. (2003): Habitat fragmentation and amphibian conservation.


7)      Pesticides

Required Readings:


a.       Be able to describe what evidence exists that wind-blown pesticides from agriculture areas are causing amphibians declines

Wells: p. 840

b.      Be able to describe how does atrazine affects amphibian reproduction.

Wells: p: 840

c.       Be able to elaborate on some of the unexpected synergistic effects of pesticides. 

Wells: p. 842

d.      Be able to describe how pesticides impact amphibian immune function.

Wells: p. 842

Supplemental readings:

1.      Wells pp. 839–843

2.      Relyea and Mills (2001): Predators make pesticides more lethal.


8)      Bd and Ranavirus

Required Readings:

·         Wells:

a.       What was the first country to report die-offs from Bd?

Wells: p. 831

b.      Be able to describe some weaknesses of the “Out of Africa” hypothesis.

Wells: p: 834

c.       Be able to describe how global climate change may be contributing to the emergence of Bd. 

Wells: p. 834

d.      Be able to describe how differences in antimicrobial peptides (AMPs) associated with the skin of amphibians may be responsible for differences in susceptibility to Bd that have been observed among amphibian species.

Wells: pp. 835-837

·         Gray et al. (2007): Ranavirus and Cattle

·         Bryan et al. (2009): Ranavirus and Disinfectants

Supplemental Readings

            Wells: pp. 831-838

            Bd Treatment


9)      Conservation

Required Readings:

·         No Required Readings

Supplemental Readings

·         Recommendations for Riparian Buffers: Salamanders (Crawford and Semlitsch 2007)

·         Recommendations for Wetlands Buffers: Amphibians and Reptiles (Semlitsch and Bodie 2003)


Podcasts: (MP3 Format)

1.      iTunes Instructions: (you must first download iTunes to Listen to Podcasts)

1)      Go to http://itunesu.utk.edu

2)      Click on "Download iTunes & Quick Time"

3)      Click on “Download iTunes Free”

4)      Save iTunes to your hard drive and install.

2.      Link to iTunes to Listen to Podcasts:  Launch Podcasts in iTunes U  


Slides: (PDF Format)

Lectures:  Test 1

                   Tennessee Anuran Identification” (PDF)            Tennessee Anuran Identification     (MHT)

                   Tadpole Identification(Nathan Haislip, UTK)

                   Tennessee Salamander Identification(Guest Lecturer: Matthew Niemiller, UTK)

                   Amphibian Phylogeny and Evolution(Jason Hoverman, UTK)

                   Global Amphibian Declines

                   Amphibian Reproductive Strategies(Guest Lecturer: Dr. Sandy Echternacht, UTK)

 Courtship and Mating

 Tadpole Development and Metamorphosis


Lectures:  Test 2

Amphibian Dissection Lab(Jason Hoverman and Nathan Haislip, UTK)

Phenotypic Plasticity(Jason Hoverman, UTK)

Community Ecology(Jason Hoverman, UTK)

Community Ecology(Nathan Haislip, UTK)

Amphibian Sampling 

Disease Sampling Techniques(Nathan Haislip, UTK)

Bd and Ranaviruses

Amphibian Pathology(Guest Lecturer: Dr. Debra Miller, UGA)

Amphibian Conservation

Graduate Student Lectures: Test 2

1)     Elizabeth Summers (Estivation and Hibernation)

2)     Wyn Miller (Landscape Ecology)

3)     Kevin Hamed (Pesticides)

Undergraduate Mini-Presentations: Test 2 (Causes of Die-offs and Declines)

(1)     Global warming (Hamlington)

(2)     Ozone depletion and UV-B radiation (Young)

(3)     Acid rain (Pace)

(4)     Silviculture (Johnson)

(5)     Fragmentation (Robinson)

(6)     Roads (Alexander)

(7)     Exploitation (Blystone)

(8)     Introduced Species (Carr)

(9)     Cattle (Spain)

(10)     Agricultural fertilizers (Dutkosky)

(11)     Pesticides (Simcox)

(12)     Endocrine disruption (Rasnick)

(13)     Batrachochytrium dendrobatidis (Moss)

(14)     Saprolegnia (Van Mater)

(15)     Ribeiroia (Reasor)


Diversity and Evolution Lecture


1.       Video 1: Parental care in caecilians.  Female caecilian “feeding” her young sloughed skin that is high in fat content.  From Life in Cold Blood.  BBC production.  Hosted by David Attenborough.  Link: http://www.bbc.co.uk/sn/tvradio/programmes/lifeincoldblood/video.shtml?licbtt03


2.       Video 2:  Red-legged salamanders mating.  Male deposits a spermatophore that is subsequently picked up by the female.  From Dr. Stevan Arnold’s website.  Dr. Arnold is at Oregon State University and studies evolutionary biology and ecology (http://oregonstate.edu/~arnoldst/).  Link: http://oregonstate.edu/~arnoldst/shermani transter.avi


3.       Video 3:  Plethodontid salamander feeding. Hydromantes platycephalus is able to project its tongue more than half its body length to capture prey.  From Dr. Stephen Deban’s website.  Dr. Deban is at the University of South Florida and studies evolutionary biology and functional morphology (http://autodax.net/index.html).  Link: http://autodax.net/hydromovie.html


4.       Video 4:  Hellbender feeding. Cryptobranchus alleganiensis uses suction feeding to consume prey within its aquatic environment. From Dr. Stephen Deban’s website.  Link: http://autodax.net/Cryptomovie.html


5.       Video 5: Bolitoglossa schizodactyla walking. Bolitoglossa schizodactyla has unique adaptations for climbing.  One of your assignments is to read about this in your book.  From Amphibia Web website..  Link: http://amphibiaweb.org/sounds/Bolitoglossa_schizodactyla.mov


6.       Video 6: How anurans jump. Video discussing the evolution of jumping in anurans. Dr. Anne Maglia’s is at the Missouri University of Science and Technology and studies amphibian development and evolution (http://web.mst.edu/~magliaa/index.html).  Video at: http://www.midwestfrogs.com/


7.       Video 7: Male poison arrow frogs wrestling over territories.  From Life in Cold Blood.  BBC production.  Hosted by David Attenborough.  Link: http://www.bbc.co.uk/sn/tvradio/programmes/lifeincoldblood/video.shtml?licbtt02


8.       Video 8: Male poison arrow frogs and their tadpoles.  Males carry their tadpoles on their backs to bromeliads to complete their larval development.  From Life in Cold Blood.  BBC production.  Hosted by David Attenborough.  Link: http://www.youtube.com/swf/l.swf?swf=http%3A//s.ytimg.com/yt/swf/cps-vfl74240.swf&video_id=64y15Ho6d84&rel=1&eurl=&iurl=http%3A//i3.ytimg.com/vi/64y15Ho6d84/hqdefault.jpg&sk=ynZ1LV0QasPhd6zqZEUWSabwnNSPShPiC&use_get_video_info=1&load_modules=1&hl=en


Amphibian Declines Lecture


1.       Worldwide Amphibian Declines: How big is the problem, what are the causes and what can be done?  Interview with Vance Vredenburg, Robert Drewes, Tyrone Hayes, and Karen Swaim.  Video at: http://amphibiaweb.org/declines/declines.html.


Courtship and Breeding Lecture


1.       A Unique Way to Travel: http://www.youtube.com/watch?v=HmLS2WXZQxU

2.       Frog-eating Bat: http://encarta.msn.com/media_461575349/frog-eating_bat.html

3.       Salamander (P. shermani) courtship (mental glands slapping and spermatophore transfer): http://plethodon.science.oregonstate.edu/behavior.html#

4.       Spotted salamander ovipositing eggs: http://www.youtube.com/watch?v=5nTm0eDDQko&feature=channel_page

5.       Wood Frogs (explosive breeders, amplexus and egg fertilization): http://www.midwestfrogs.com/

6.       Pleurodema diplolister (Foam nest construction, amplexus and egg fertilization): http://amphibiaweb.org/sounds/Pleurodema_diplolister2.mov

7.       Hellbender Videos (external fertilization, Video III): http://www.hellbenders.org/hellbendervideo.html


Tadpole Development Lecture


1.       Time-lapsed (11 d) Embryo Development (Rana temporaria):  http://en.wikipedia.org/wiki/File:Frog_spawn_time-lapse.gif

2.       Tadpoles Scavenging Dead Frog: http://www.youtube.com/watch?v=KXH6YPWlQaI&feature=PlayList&p=9D79119137C269D7&index=0&playnext=1

3.       Tadpole Schooling and Bobbing Behavior (Leptodactylus insularum): http://www.ristohurme.com/insularum.htm

4.       Tadpole Development for Archey’s Frog (Leiopelma archeyi): http://www.arkive.org/archeys-frog/leiopelma-archeyi/video-09.html

5.       Tadpole Metamorphosis (Spotted tree frog, Litoria spenceri): http://www.arkive.org/spotted-tree-frog/litoria-spenceri/video-09.html



TWRA Amphibian Identification:      http://www.state.tn.us/twra/tamp/frogs.html 

LEAPS Anuran Identification:                       http://www.leaps.ms/Tn.%20Frogs%20ID%20Page.htm


Video Clips of Frog Calls:                  http://www.midwestfrogs.com/


TWRA Salamander Identification:     http://www.state.tn.us/twra/tamp/salamanders.htm


Amphibians of the Great Smoky Mountain National Park (Dodd 2003):            http://fisc.er.usgs.gov/c1258_Dodd/


Southeast Tadpole Identification:            http://fl.biology.usgs.gov/armi/Guide_to_Tadpoles/guide_to_tadpoles.html


ARMI 5-year Report (Amphibian Declines):            http://www.fort.usgs.gov/products/publications/21733/21733.pdf


Global Amphibian Assessment:                      http://www.globalamphibians.org


USGS Field Guide to Malformations of Frogs and Toads:            http://www.nwhc.usgs.gov/publications/fact_sheets/pdfs/frog.pdf


PARC Habitat Management Guidelines for Amphibians and Reptiles of the Southeastern United States: http://www.parcplace.org/habitat_management_guide.html


Previous WFS 433/533 Websites

Spring 2008:  http://fwf.ag.utk.edu/mgray/wfs493/493home2008.htm

Spring 2007:  http://fwf.ag.utk.edu/mgray/wfs493/493home2007.htm